1. Centre for Biological Diversity and Scottish Oceans Institute School of Biology University of St Andrews St Andrews FifeKY16 9TH UK
2. Department of Biology University of Vermont Burlington Vermont05405 USA
3. Department of Mathematical Sciences Loughborough University Loughborough LeicestershireLE11 3TU UK
4. School of Biology and Ecology Sustainability Solutions Initiative University of Maine Orono ME04469 USA